Superior growth process of creeping ramets of Phragmites australis and its physiological mechanisms in an alkaline meadow in Northeast China

doi:10.17521/cjpe.2023.0158

Abstract

Abstract:

Aims The reed (Phragmites australis) is a long-rhizomed clonal plant, which is distributed worldwide and has the plasticity and adaptability to change its morphology, even growth form with environmental change. The creeping ramets of reeds are a special growth form generated from their rhizomes extending out of alkaline soil patches. This study aims to explore the growth of creeping ramets and their underlying mechanisms.

Methods Using methods such as regular tracking of ramet growth by hanging tags, measurement of photosynthetic physiology of leaves of different ages, and determination of ¹⁵N isotope transfer, we measured and analyzed the growth rhythms and patterns of creeping reed ramets, their photosynthetic characteristics, and indicators of physiological integration between ramets.

Important findings We found that in highly alkaline areas, reeds’ creeping ramets displayed different growth patterns compared to control, upright ramets. After 120 days of growth, the creeping ramets had an average length of (685.25 ± 118.75) cm, and their average growth rate during the observation period was (6.64 ± 3.51) cm·d^-1. This was 15.4 times faster than the control, upright ramets, indicating a logarithmic allometry growth process that started quickly but then slowed down. In contrast, the control ramets showed a relatively stable linear isogonic growth process. Young leaves at the top of the creeping ramets had the same maximum photosynthetic capacity as mature functional leaves. The net photosynthetic rate of leaves on the creeping ramets varied in a logistic curve with increasing leaf order, while the control ramets varied in a quadratic curve that first increased and then decreased. Moreover, the theoretical maximum net photosynthetic rate of creeping ramets was 19.4% higher than that of the control ramets. Creeping ramets treated with ¹⁵N isotopes had significantly higher levels of ¹⁵N abundance in various organs compared to untreated creeping ramets. Creeping ramets are a new adaptive feature of this widespread plant in extremely harsh alkaline habitat patches. The superior growth of creeping ramets is attributed to the high photosynthetic rate of young apical leaves and the physiological integration between tufted basal ramets and creeping ramets. This study provides new insights into the adaptation of reeds to extreme habitats and offers a method for analyzing the superior growth of creeping ramets based on matter production and physiological integration, with important theoretical implications.

Key words: clonal plant, physiological integration, ¹⁵N isotope, ramet growth form, assimilate transfer, extreme habitat, plant ecology

HAN Da-Yong, LI Hai-Yan, ZHANG Wei, YANG Yun-Fei. Superior growth process of creeping ramets of Phragmites australis and its physiological mechanisms in an alkaline meadow in Northeast China[J]. Chin J Plant Ecol, 2025, 49(2): 320-330.

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URL: https://www.plant-ecology.com/EN/10.17521/cjpe.2023.0158

https://www.plant-ecology.com/EN/Y2025/V49/I2/320

Figures/Tables 9

Fig. 1 Bare patches (A) and composition of the tufted Phragmites australis growth form (B) in fenced plots in a salinized Leymus chinensis meadow during mid-July. CR, creeping ramet; LR, lying ramet; NR, nodal ramet; OR, oblique ramet; UR, upright ramet.

Table 1 Basic statistics on the growth rate of the creeping ramets of Phragmites australis at different time periods (cm·d-1)

时间段(月-日) Time period (month-day)	样本数 n	最大值 Max	最小值 Min	平均值 Mean	标准差 SD	变异系数 CV (%)
4-25-6-04	7	4.20	1.26	2.94^c	1.12	38.0
6-05-6-14	7	15.72	2.09	8.59^ab	4.72	54.9
6-15-6-24	7	15.08	7.92	11.46^a	2.67	23.3
6-25-7-04	7	16.22	8.21	10.74^a	2.78	25.9
7-05-7-14	7	12.95	5.27	8.51^ab	2.88	33.9
7-15-7-24	7	10.94	1.70	6.06^b	3.50	57.8
7-25-8-04	7	11.06	1.70	6.66^b	4.41	66.2
8-05-8-14	7	8.39	0.33	3.17^bc	2.75	86.6
8-15-8-24	7	6.71	0.04	1.64^c	2.29	139.8

Fig. 2 Comparison of differences in growth rate (mean ± SD) between the creeping ramets and the control upright ramets of Phragmites australis at different time periods. *, significant difference (p < 0.05) between different ramets at the same period; ns, no significant difference (p > 0.05).

Table 2 Parameters and significance tests of the fitting relationships between the length (y, cm) of the creeping (CR) and control upright ramets (CK), and growth time (x, d) of Phragmites australis

分株 Ramet	样本号 No.	样本数 n	函数类型 Function type	方程参数 Equation parameters		R²	p
分株 Ramet	样本号 No.	样本数 n	函数类型 Function type	a	b	R²	p
匍匐型 CR	1	9	对数 Logarithmic	-1 322.8	378.34	0.92	<0.01
	2	9	对数 Logarithmic	-1 686.9	502.44	0.99	<0.01
	3	9	对数 Logarithmic	-1 727.7	522.14	0.93	<0.01
	4	9	对数 Logarithmic	-2 316.3	635.67	0.98	<0.01
	5	9	对数 Logarithmic	-1 682.3	514.24	0.95	<0.01
	6	9	线性 Linear	-238.1	8.33	0.99	<0.01
	7	9	对数 Logarithmic	-2 577.9	709.60	0.96	<0.01
	平均值 Mean	9	对数 Logarithmic	-1 946.3	556.36	0.99	<0.01
直立型对照 CK	1	9	线性 Linear	17.80	0.27	0.99	<0.01
	2	9	对数 Logarithmic	-57.88	23.10	0.98	<0.01
	3	9	线性 Linear	23.25	0.32	0.94	<0.01
	4	9	线性 Linear	13.50	0.44	0.99	<0.01
	5	9	线性 Linear	24.51	0.37	0.99	<0.01
	6	9	线性 Linear	17.38	0.45	0.99	<0.01
	7	9	对数 Logarithmic	-97.72	35.43	0.99	<0.01
	平均值 Mean	9	线性 Linear	0.38	19.18	0.97	<0.01

Fig. 3 Observed values and fitting curves on the relationship between the growth time and the average length of creeping ramets (A) and control upright ramets (B) of Phragmites australis (mean ± SD).

Fig. 4 Comparison of photosynthetic rate (Pn) and transpiration rate (Tr) (mean ± SD) of leaves at different orders of creeping Phragmites australis. Different lowercase letters indicate significant difference (p < 0.05) between leaf orders.

Fig. 5 Observed values and fitting curves of photosynthetic rate (Pn) and transpiration rate (Tr) of Phragmites australis leaves with varying leaf order on creeping ramets (A1, B1), nodal ramets (A2, B2) and control upright ramets (A3, B3).

Table 3 Parameters and significance tests of the relationships between the photosynthetic physiological indices (y) and leaf order (x) of Phragmites australis creeping ramets (CR), nodal ramets (NR) and control upright ramets (CK)

因变量 Dependent variable (y)	分株 Ramet	样本数 n	方程参数 Equation parameter				R²	p
因变量 Dependent variable (y)	分株 Ramet	样本数 n	A₁/a	A₂/b	x₀/c	m		p
光合速率 P_n (μmol·m^-2·s^-1)	匍匐型分株 CR	15	14.40	22.83	11.62	10.21	0.96	<0.01
	节生型分株 NR	8	2.89	4.81	-0.35		0.88	<0.01
	直立型分株 CK	8	-8.50	6.59	-0.40		0.94	<0.01
蒸腾速率 T_r (µmol·m^-2·s^-1)	匍匐型分株 CR	15	5.04	7.21	12.09	15.63	0.94	<0.01
	节生型分株 NR	8	1.33	2.18	-0.19		0.93	<0.01
	直立型分株 CK	8	-4.71	2.90	-0.19		0.92	<0.01

Fig. 6 Comparison of 15N abundance between tracer-treated and control Phragmites australis creeping ramets in different organs and sections (A), and among tracer-treated organs (B) (mean ± SD). L, leaf; LS, leaf sheath; S, stem. 1, 2 and 3 represent upper, middle and lower sections. *, significant difference (p < 0.05) between tracer treated (15N) and control treatments. Different lowercase letters indicate significant difference (p < 0.05) among organs.

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